植物科学最前線

6: 2 ( 2015)"3&JN>E1"+'0$(3BE.46#BEļǜ Ă, ǋÉ Ʈĥ, Ɵƞ ŤĥǪãħȏȝũǬęʍǗȖħȏȝ\_230-0051 ȔĢľȇƺǑŀ˘ɵäƝņǲ 1-7-22
Molecular mechanisms on callus formation: Accelerators and Brakes
Key words: callus, dedifferentiation, phytohormone
Akira Iwase, Momoko Ikeuchi, Keiko Sugimoto
RIKEN Center for Sustainable Resource Science
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A. Iwase, M. Ikeuchi & K. Sugimoto-1
BSJ- Revi ew 6: 2 ( 2015)

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6: 3 ( 2015)

ōť*Îĥ~Mi[})" $, ōť¶ʤĈĥ˚FQ\˛&ŭÕĈĥ˚tO˛)Î$ȶ
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A. Iwase, M. Ikeuchi & K. Sugimoto-2
BSJ- Revi ew 6: 3 ( 2015)

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6: 4 ( 2015)

ZʂĳŞ*Ĉĥ˚² ,ǏŞʱȳȚ˛, Ïƈ˅ =*ēĐťÎ*³Ȼ¶ʤ, ¡*ȸɆ(' =ƣ?ÉǮ*ʈĈĥ˚ưǣyȦ˛*ƽɤȦ, Ʒ(ɴĈŋʵ)(!$ ?&Ɍ =@?4¿ĬʂĳŞ*uR |RĈĥ*ƼɑǧŔ </Ƽɑƽžưǣ­%+, ȸɆē˓ȱ
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2011a), ƭ*ŅȵɐȿŲ)ʻ?ʭ¨ĥ˚² , PLETHORA1˛CǾǩ$ ?MZ9, ɢˇÎɯȸɆ*ŅȵɐȿŲ)Śɴ(ʭ¨ĥ(² , WUSHEL 9 SHOOT MERISTEMLESS)CǾǩ$?8*ɸ=@$ >, MZ*ɵȄ8ÊÎã*Àù8Ƕ(!$ ?īˀ, PLT1 CǾǩ$?MZ+, ưǣyCû4( ēĐ)MZCȘư?&ƭCÊÎã?Àù˄ł)˗  *< (ʭ¨ĥǾǩusEH*Ƕ(!MZ+, ÷*ȸɆǢC÷*ēĐƢ¥%ē˓ˀ)8Íǩ ?XHjh[h*ȸɆē˓ȱ%ɝǰ =@? Callus Inducing Medium(CIM; Valvekens et al., 1988)%ē˓ƭ*ȸɆǢ%+, ¿Ĭʮ«*5(=˄¿Ĭʮ« =8M
 Z  Í ǩ  $  ?  Sugimoto = + ,  * ȱ * ˄ ¿ Ĭ ʮ «=*MZōťƔ),SCARECROW(SCR)9 WUSHEL RELATED HOMEOBOX5 (WOX5)('ƭ*Ņȵɐōť)ʻ?{Mʭ¨ĥŌǾǩ?&CǾɵ$ ?4, »ƭ°=@((?ĝǶ­%+M
Zōť@((?& =, *Ƣ¥*MZ+»ƭêĔōť*ȹʒCȹǱ$ōť@$ ?&Cĕü$ ?(Sugimoto et al., 2010)Ɗ, ȕʩɸį$ ?¿Ĭʮ«*MZ%+@=*{Mʭ¨ĥ*Ǿǩ+ɸ=@( 4, »ƭ°=@((?ĝǶ­%8, ¿Ĭʮ«%+MZãʏ?@=*ɸį =, ¿Ĭʮ«%°=@$ ?MZ+, ƭ*{Mʭ¨ĥCǾǩ?˄¿Ĭʮ«*MZ&+, ĵ(&8?șň+Ƕ(?ȹʒ%°=@, Ƕ(?ʭ¨ĥǾǩusEH*MZ%?&Ə= &(!(Iwase et al., 2011a)*< )MZ&

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6: 5 ( 2015)

1932)4ȁÙ@ƻƛƻȁCÊǮ?ǩʋ+, Ȳ 200 ń×)+ʀȌ@ȏȝ@$ &ųſ@$ ? (Stobbe et al. 2002)*ʧș*%+, surface callus &ÿ,@?ȵɐėōť
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BSJ- Revi ew 6: 6 ( 2015)

植物科学最前線

6: 7 ( 2015)ȁ9˖)ĉ4@$ƛʮ9Łʮ)ªȸɆ (psuedophloem)9, 4ÎɯȸɆɸį@?(Lee,1955)÷Ʒ)Ȥ 3 ɉ*IGZ%? Rice gall dwarf virus +, Hk9U}P('*HkȖưǣC
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B:?ɪ0C"=?Ǝɪˏ (Tooker et al., 2008)ȦŴ=@?

@=*)+, °ǣ)ǭĠ(ɮĬCʏ8*?, Ţƨ9ǸŘǩ@?ˀ*Îĥ~Mi[}CɹƏ?+ʘƳȀ)8Ġĝʲɴ%?Ɗ, ưǣ)&!$8Ŧˏ)&!$8˄ł)ʲɴ(Ǻ?*"+, ƭȭɣ{~Ȗưǣ)°=?ƭȭ%?, @)ʻ$+,ıĽ=)<?Ȥ 6 ȢCìǡ@ 

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ưǣ*?Țʺ%˂C?&, ʁǮ˂Ț*¹­)MZ%?&?@+ʭ¨Ȁɖǹ&ÿ,@$ >, Ft hȖFt hĺ, hZȖaI\FWNLĺ, hZȖiUaF
hĺ, Ȗĺ('%ĕü@$? (Ahuja, 1965) ²,, _nU*˂Ț Nicotiana

glauca× N. langsdorfii )°=@MZ+ưǣyCǓÞ( ēĐ%8Ƚ£ē˓óɑ%>, 4¹­CÊǮ?8%?(White, 1939; Ichikawa and Syōno, 1988)˅ǿ &), *_nU*ʭ¨Ȁɖǹ%+¹­*ɋãʤ6&MZōťʤ6ɡ ¹­%8, ưǣ­)¿C"?&%MZōť@?(Udagawa et al., 2004)êĈCȝƏ?ʢ*ȏȝ =, ¼Ǡ,N. glauca *Tl})ǆǥƭCưǣ)°=?&%Ƙø(nQdF Agrobacterium rizogenes *root-inducing (Ri)u Z|g*ʭ¨ĥʯÐ)<ªˊĒɵ" >, īˀ*ˊĒ)Ug@ʭ¨ĥ+_nU)ōţĝãCʏƼɑ?&Î !(Aoki and Syono, 1999) (=, _nU%+*ʭ¨ĥʯÐCŲ( Ț*ȸ5öB%8ʭ¨Ȁɖǹʏ?&B>, ǈŃ¨ƀ)<!$ŵÅ@&Ɍ =@?*nQdFǱƣ*ʭ¨ĥɖǹã*?
êĈ%+( &ĕü@$ ?(Kung, 1989)ưǣy&*ʻµ)" $+ÉǮʴ*ĝãʏ$ ?&

ĕü?(Kehr, 1951; Kung, 1989; Ichikawa and Syōno, 1991), Îĥw

%*ɹƤ+4>ʤD% ( p{*Țʺ˂Ț Helianthus annuus × H. tuberosus )ɸ=@?
ʭ¨Ȁɖǹ%+, ɏǾǮ9ÎɯȸɆ*ōť)ʲɴ(ʭ¨ĥ*ǶũȀ(Ǿǩĕü@$?

(Chiappetta et al., 2006, 2009)

3. "3&JN>E1"+'0 ¿ĬʂĳŞ*MZōť+¿ò%ǤǶȀ)ɸ=@?,@+ʮ«ǤǶȀ(ÕŕƼƶĦď?&CȓĄ$ ?4, ưǣȵɐ%+ňÎãȵɐ 1 " =%8MZãCȹ$¹­
CÊǮ%, ÎãÆɑŞCƘ?& 1950 ń£őç1970 ń£) $Ƌ)Ə= )@$?˚Steward 1959, Nagata and Takebe 1971˛, ʠł*ǾǮˡÎãǄʿ%+*ɑÝCŭ ʚ6ƼƶŚɴ)ʪ ( æ¯ń)Ǖ?ưǣ*Îĥʭ¨ħɹƤ*ʤĹ)©!$, MZã)ʻ
?Ʒ(ĝǶ­è˃@$@)<>, MZã)ʻ?ʭ¨ĥ8Œ)Ə=A. Iwase, M. Ikeuchi & K. Sugimoto-6
BSJ- Revi ew 6: 7 ( 2015)

植物科学最前線

6: 8 ( 2015))(!$$ ? (ɭ 1)@=*Ĉĥ+, MZãC¶ʤ?FQ\Ĉĥ&, ŭÕ?tOĈĥ)ĠÓ%?īˀ, ȕʩ*ȏȝ =8¿ò%Ǿǩ¶ʤ, MZãC¶ʤ?ʔËĈĥ(Iwase et al. 2011a, 2011b)ɵ" >, =)ƗȷÎãȵɐɕÎã( < )?Ƽƶ
˚Ikeuchi and Iwase et al., submitted˛8ɵ ""?*Ȫ%+, @=*ĈĥCȶ?&&8), Ǥ)MZã*ǤŘ*"%?ȵɐÎɯ*ƍʤ9Êʹ&*ʻʢ)ȋȄ$Ƶʄ  aw[jhchQOXPLW]ubTZ_du¡~RovG^¡~
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A. Iwase, M. Ikeuchi & K. Sugimoto-7
BSJ- Revi ew 6: 8 ( 2015)

植物科学最前線

6: 9 ( 2015)

3-1. SX/325qn"3&@$(3/.46#BE 2-1 Ȫ%ɺ@< ), CIM %ōť@?MZ)+»ƭêĔōťCȹǱ8*?
(Sugimoto et al. 2010)»ƭêĔōť+LOX*XRh)<!$ǏŞã@?, *ȹʒ%Ƽɑ?&Ȍ=@$ ? LATERAL ORGAN BOUNDARIES DOMAIN (LBD) sE|ʔËĈĥ)ĺ LBD16, LBD17, LBD18, LBD29 *ʭ¨ĥ+, @@XHjh[h%ʧÚǾǩ?&ưǣyCǓÞ( ēĐ)$8MZCōť?(ɭ 1; Ċ 3)4, Ƽɑƽž(lbd16)4+ŭÕđ(35S:LBD16-SRDX)*ưǣ­%+, ʞ)ưǣyCû6ēĐ(CIM))$8MZãŭÕ@?(Fan et al. 2012)LOX)śȨ LBD *ǾǩCǀ)Õŕ?
ʔËĈĥ&$ AUXIN RESPONSIVE FACTOR7 (ARF7) & ARF19 ĕü@$ ?(Okushima
et al., 2007; Lee et al., 2009), arf7 arf19 ʲĝǶ­%8MZã+ŭÕ@?=) arf7 arf19 ư
ǣ% LBD16 CʧÚǾǩ?&Ê/MZãɑćŖ? =(Fan et al. 2012), ARF7 9 ARF19*ǐĈĥ&$Ħď? LBD ʔËĈĥɉ, »ƭêĔōťCȹǱMZōť*FQ\Ĉĥ%?&+Ə= %?ȥɍ=8 LBD16 CʧÚǾǩXHjh[hưǣ­%, ưǣyCû4( ēĐ)$8ƭ)MZōť@?Cȑʀ$ ? (Ikeuchi et al.,2013˞Ċ 3)LBD &ȵɐýƚÊʹ*ʻʢ&$+, LBD18 & LBD33 *ʴ­ E2 PROMOTER
BINDING FACTOR a (E2Fa)*ǾǩC˗7?&

ĕü?(Berckmans et al., 2011)ʔËĈĥ

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QLOX)<?ȵɐýƚÕŕ*"CŰ!$ ?&Ɍ =@? ȵɐýƚCÊǏŞã?)+, ȵɐýƚ*tOCĞ&

ŧǴ8?Cyclin Dependent

Kinase (CDK)CʼĬ? KIP-RELATED PROTEIN (KRP)CUg?ʭ¨ĥ+LOX)<!
$ǾǩŭÕ@?ʔË*F`u__oQʎ%? PROPORZ1 (PRZ1)*u\Z)ʻ
$ ?ĕü@$ ?(Anzola et al., 2010˞ɭ 1) prz1 ĝǶ­+, ʳǮƫ»ƭCōť?LOXǛň*ēĐ%8MZCōť?*ʧÚ(ȵɐÎɯ+ KRP2, KRP3, KRP7 ʭ¨ĥ*ʔËʴ¬ &)<!$ŋʏ@$ ?(Sieberer et al., 2003)PRZ1 + KRP2, KRP3,
KRP7 ʭ¨ĥ*u_)@@ȅŸȺö?&9, prz1 ĝǶ­%+ KRP7 * 5’UTR ˊĒ)Å?pZf H3-K9/K14 *F\aã{Q*w?& =, LOXÌǪ)<!$pZf*F\aãw?&% KRP *Ǿǩʴ>, ȺƦ&$ʧÚ(ȵɐÎɯŋʏ@$ ?&Ɍ
=@$ ?(Anzola et al., 2010)īˀ
KRP ǾǩʴCFa\ZǍ%ŭ
?&MZã¶ʤ, 4 prz1 ĝǶ­% KRP7 *ǾǩʴC?&MZã*ōʎŭ =@?&8ĕü@$ ?(Anzola et al., 2010)LOX)<!$ PRZ1 '*< )Õŕ@$?

+Əȑ)(!$(

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@=*ȺƦ+LOX)<?ȵɐÎɯǏŞãȹʒ)+, PRZ1 ´ĦȀ(A. Iwase, M. Ikeuchi & K. Sugimoto-8
BSJ- Revi ew 6: 9 ( 2015)

植物科学最前線

Q{aÕŕC KRP *ʭ¨ĥǾǩŭÕ&6: 10 ( 2015)ʨȧ?&CȓĄ?8*%?

 WHfMHiʻʢĈĥ%MZōť&*ʻƏ=)(!$?8*)+, type-B

ARABIDOPSIS RESPONSE REGULATORs (ARRs)?(ɭ 1; Ċ 4)type-B ARR + B:?ťÎÕŕȱ)<!$ʱã@ǏŞã?ʔËĈĥ%>, ğ*ǐʭ¨ĥ*ǾǩCʂĳ
?(Hwang et al., 2012)WHfMHiCû6ēĐ% ARR1 CǾǩXHjh[hCɎ$?
&, MZōť¶ʤ?(Sakai et al., 2001)4ʱãg~HCƽžşłȀǏŞđ)
 ARR1 9 ARR21 ÎĥCŌÕǾǩXHjh[h%+, ưǣyCû4( ēĐ%8
MZCōť?(Sakai et al., 2001; Tajima et al., 2004)*& =, ARR1 ´ĦȀ(WHfMHiśȨȱ*ǏŞã8MZōť)+æÎ%?&Î ? ȵɐÎɯ*ÊǏŞã)ʻ? type-B ARRs *_Tbfºɰ&$+, Cyclin D3 (CYCD3)
ƘÝ 8@( CYCD3 +WHfMHiÌǪőƔʺ%Ǿǩʴ¶ʤ?), CYCD3 *ʧÚǾǩ+WHfMHiCû4( ʂĳēĐ)$MZãC¶ʤ?(Riou-Khamlichi et al.,1999)=)˅ǿ ), CYCD3;1 &*yR%? CYCD3;2  CYCD3;3 *ʲĝǶ­%+WHfMHiśȨŭÕ@?& =, CYCD3 +WHfMHiXRh*ǐĈĥ&$Ƽɑ$ ?ĕü@$ ?(Dewitte et al., 2007)
AP2/ERF ʔËĈĥsE|)ĺ ENHANCER OF SHOOT REGENERATION1 (ESR1)& ESR2+, WHfMHiCMZã)ʻ?¡*ºɰĈĥ%?(ɭ 1; Ċ 4)ESR1 8 ESR2
8XHjh[h)$èǦ%ʧÚǾǩ?&%, ưǣyCû4( ēĐ%8M

ZǮ?(Banno et al., 2001; Ikeda et al., 2006)4 BOLITA (BOL)&

FQawX_R

H%+, ESR2 *ʧÚǾǩ $ >, %8MZãɸį@$ ?(Marsch-Martinez
et al., 2006)ESR CʧÚǾǩưǣ%+WHfMHiśȨƍʤ$ >, 4, WHfM
Hi\u_*ƼɑƽžĝǶƫ%? cytokinin response1/Arabidopsis histidine kinase4 % ESRCǾǩ?&WHfMHiśȨ*"*ųƸ%?ɢɥ*ÊǮɑŨ?(Banno et al., 2001;
Ikeda et al., 2006)ESR2 + CYCD1;1 & DOF ʔËĈĥ*"%? OBF BINDING PROTEIN1(OBP1)*ǾǩCȅŸʂĳ?&ĕü@$ ? (Ikeda et al., 2006)* OBP1 +ʧÚǾǩ)<!$ȵɐýƚʻʢʭ¨ĥ*ǾǩCʂĳ, G1 ƚCȍ?%ȵɐýƚC¶ʤ$ ?&ĕü@$ ?(Skirycz et al., 2008)È­Ȁ)+ OBP1  CYCD3;3 & S ƚǤǶȀ(ʔËĈĥ%
? DOF2;3 *u_)ȅŸȺö?&ȓ
@$ ?(Skirycz et al., 2008)MZã*ˀ)@=* ESR CʔËkbfQīˀ)ȵɐýƚCǏŞã?* )" $+Ɩ(?ƲɽŚ
ɴ%? , @=*Ȍɵ+, ȵɐýƚ*ÊǏŞãƷ(ʿĻ*ʔËĈĥ)<!$Ƃʯ@$?óɑŞCȓ$ ?4, 88 ESR1 +ʧÚǾǩ)<!$WHfMHi˄´ĦȀ)ɢɥÊǮCʏʭ¨ĥ&$è˃@$ >(Banno et al.
2001), MZã&ɢɥÊǮ&*ʻʢCɹƏ 
7*ʲɴĈĥ&$8ǎȄ@?

A. Iwase, M. Ikeuchi & K. Sugimoto-9
BSJ- Revi ew 6: 10 ( 2015)

植物科学最前線

6: 11 ( 2015)

3-2. =FLaM"3&@$(3BE ¿Ĭ)<?ȵɐuR |RǩʋğȵɐǮǣ)ņɸ=@?&+ 2-2 Ȫ%ʜ2ư
ǣ%ɸ=@?MZã)$8, ɛǠǳ ȸɆē˓Ƣ¥˚2-1 Ȫ˛ )ʻB=, ¿Ĭʲɴ(ŋʵ)(!$ ?*ȹʒ)ʻB?ÎĥƼƶ+, ʛń)(!$ǚ " *ʲɴĈĥè˃@$)( ȕʩǩďȏȝCʤ7$? AP2/ERF s E |   * ʔ Ë Ĉ ĥ WOUND INDUCEDDEDIFFERENTIATION1 (WIND1)+, XHjh[h*MZǱƣ*ē˓ȵɐ%˗Ǿǩ$?Ĉĥ&$ʬů@$(Iwase et al. 2005)WIND1 &*yR%? WIND2, WIND3,
WIND4 C@@XHjh[h%ʧÚǾǩ?&, ưǣyCû4( ēĐ%8ɢɥ,ɏʕ, ƭ(')MZǮ?(ɭ 1)4ʧÚǾǩ­ =Ŕ=@MZ+ysēĐ%Ƚ£ē˓óɑ%? (Iwase et al. 2011a; 2011b)XHjh[h*ʛɃȚ%? Thellungiellahalophile * WIND1 yR(ThWIND1-Like)CXHjh[h%Ǿǩ?&, 9+>ĞĈŞ*y˄´ĦȀ)MZōť@?(Zhou et al., 2012)4XHjh[h WIND1(AtWIND1)*MZʂĳɑ+, h_k, f{f, _nU%8ȑʀ@& =, WIND1 Îĥ)<?MZãȹʒ+ĵ(&8?ȫĉ*íĥɥưǣ%+·Ħ@$ ?< %?(Iwase et al. 2013)
WIND1 + RAP2.4 &8ÿ,@$ > (Okamuro et al., 1997), ¿ĬśȨŞ?&8ĕü@$
(Delessert et al., 2004)īˀXHjh[h)ò%Ǿǩʂĳ@, ¿Ĭʮ«)$ WIND1~4 +¿ĬÌǪőƅƔʺ¤É)¿

?MZãCǀ)Õŕ?&, ƼɑǧŔđĝǶ­(35S:WIND1)&ƼɑŭÕđĝǶ­(ProWIND1:WIND1-SRDX) Cǰ

ɹƤ

=Ə=

&(!$

?

(Iwase et al. 2011a)Þ $@=*ĝǶ­Cǰ ī˔ =, ȸɆē˓ȱ) ?MZōť&
Ćĩ*ÊÎã)ʻ$8 WIND ʔËĈĥʲɴ(ÁCŰ!$ ?&Ɨʛ*ɹƤ =8ɵ
$$ ?(Iwase et al. Submitted)
 WIND1 )<?MZʂĳ+, arr1 arr12 ʲĝǶ­%+ŌŭÕ@?(Iwase et al., 2011a)4
, ¿ĬZfZ+¿ò%* type-B ARR ´ĦȀ(WHfMHiśȨC˗7?, ƼɑŭÕđĝǶ­(ProWIND1:WIND1-SRDX)ưǣ­%+@ŭ =@?& =, WIND ʔËĈĥ+WHfMHiśȨC˗7$ ?&ȓĄ@$ ?(Iwase et al., 2011a)WIND1 *ǐĈĥ*ɹƤCǩďʤ7$?, ĵ(&8

type-B ARR ʭ¨ĥ*Ǿǩʴ+3&D'ĝã$(7(Iwase et al., in preparation),
type-B ARR *_oQ¸˒w%*Õŕ9 type-B ARR *cis ʯÐ)Ⱥö?¡*Ĉĥ)<?Õŕ?* 8@(  p~cNkVS%+, ɢɥ­CÏƈ?&Ïƈ˅)$ȵɐ*uR |Rʏ>, êȰ­*ˇȣŅȵɐÊǮ
$?, *ȱCǰ $¿A. Iwase, M. Ikeuchi & K. Sugimoto-10
BSJ- Revi ew 6: 11 ( 2015)

植物科学最前線

6: 12 ( 2015)ĬʂĳŞ*uR |R)ʻB?ʲɴĈĥ*ŷȴ9, ȵɐýƚÊʹ*Îĥ~Mi[}*ɹƤ8ȮÝȀ)ʤ7=@$ ?(Ishikawa et al., 2011)ɿȵ+Ȏľ)<?Ȥ 4 ȢCìǡ@  ¿Ĭ+ưǣ)$8ȸɆ9Ćĩ*ÊǮC© &ğ , *Îĥ~Mi[})" $84Î =( ğ XHjh[hȦCǰ ɹƤ%ɵ $ȸɆ9ǾǮǄʿ)<?śȨŞ*ʪ 9ʻ?Ĉĥ)ʻ$+, ǋÉ=)<?Ȥ 2 ȢCìǡ@ 4, XHjh[hɞɢ*Ǽöǩʋ)ʻB?Ĉĥ)ʻ$+, ƙǅĢ=)<?Ȥ 3 ȢCìǡ@ 

3-3. f^[, >iceHbgdOq8 BE"3&@<o  ɏŞ9ÎɯȸɆ*ƜÎãŞ*ȿŲ)ʻ?Ĉĥ*ʧÚǾǩ­, ưǣȚCąBMZC
ōť?&ĕüʛńğ5=@$ ?(ɭ 1)@+, ưǣ%ȵɐėCōť?)+ǅʖȀƜÎã(ȵɐǥţCɶĪ?ĈĥCȵɐ%ğÍ&%æÎ%?&9, ʞ)ʠł*ȵ
ɐÎã%+@=*Ĉĥ*ǾǩCƔʺȀ, ȞʺȀ)ǀÕŕ?&Śɴ%?&Cȓ$ ?CCAAT-box ȺöʔËĈĥ%? LEAFY COTYLEDON1 (LEC1), B3 g~HʔËĈĥ* LEC2, MADS box ʔËĈĥ AGAMOUS-LIKE15 (AGL15) +@@ʔËǏŞãĈĥ&$ɏǾǮƔ)Ƽɑ?, @=CèǦ%ʧÚǾǩ?&ưǣyCû4( ēĐ%8 B:? embryonic callus Ǯ?4ʭ¨ĥǾǩ*ʂĳȱCǰ $ embryonic callus Cưǣ­)°=ő)ǾǩʂĳCŭ ?%, ưǣ­CÊǮ?8%? (Lotan et al., 1998; Stone et al.,
2001; Harding et al., 2003; Gaj et al., 2005; Umehara et al., 2007; Thakare et al., 2008)h_k(Brassica
napus; Bn)%ƗÑ)è˃@ AP2/ERF ʔËĈĥsE| )ĺ BABY BOOM (BBM)8ɏǾǮƔ)Ǿǩ$?, *Ĉĥ(BnBBM)CǾǩh_k9XHjh[h, =)+UX
I, _nU, zu *ʛɃȚȦ)$8ưǣyCǓÞ( ēĐ% embryonic callus Ǯ?7, Ŕ=@Īɏ =*ưǣ­ÊǮCʠ$ưǣ­*Ěǯ1*śǰɾ5=@$ ?
(Boutilier et al., 2002; Srinivasan et al., 2007; Deng et al., 2009; Heidmann et al. 2011)@+ BBM )<? embryonic callus ʂĳ/ɏŞǧŔƼɑ, ĵ(&8?ȫĉ*íĥɥưǣ%·Ħ@$ ?
&CȓĄ$?XHjh[h)$ BBM &ʯÐ*ʛ

AINTEGUMENTA-LIKE5 (AIL5)9 PLETHORA5 (PLT5)&EMBRYOMAKER (EMK)+

ø×%8Ȍ=@$ ?, ʧÚǾ

ǩ%÷Ʒ*ǩʋʏ?&ĕü@$ ? (Tsuwamoto et al., 2010) 
RKD (RWP-RK domain-containing)ʔËĈĥ+,ˁŞʯ¼ĥ(éȵɐ)ōť9Ñƚ*ɏǾǮ%Ƽɑ
?&ĕü@$ ?RKD1 & RKD2 +éȵɐ%Ǿǩ$ ?, XHjh[h*ʧÚǾǩ­+ưǣys*ēĐ%8M
ZCōť?(Kőszegi et al., 2011)˅ǿ ),
RKD2 %ʂĳMZ*ʭ¨ĥǾǩusE
HC5?&, LOXCǰ $ʂĳM
Z<>8éȵɐ*usEH)ʛ
(Kőszegi et al., 2011)ˠ@+­ȵɐ)éȵɐƷ*ʭ¨ĥǾǩusEHCŲ$8MZCǮ=@?&CȓĄ$ >, ×ʜ
A. Iwase, M. Ikeuchi & K. Sugimoto-11
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植物科学最前線

6: 13 ( 2015)

< )MZ& !$8Ʒ(ǮǪǥţĦď?&Cšþ$ ?RKD4 +Ñƚɏ%Ǿ
ǩɏǾǮ%Ƽɑ$ ?, ʭ¨ĥǾǩʂĳȱCǰ $ RKD4 CʧÚǾǩ?&ưǣys*ēĐ%8ɥ9ƭMZã?*ˀ, Ñƚɏ%Ǿǩ$ ?ʭ¨ĥƅğǾǩ
$ >, =), MZ)(!ȵɐ% RKD4 *ǾǩʂĳCƿ7?&Īɏōťʏ?(Wakiet al., 2011)
ưǣ*ÎɯȸɆ+, ưǣ*­C°>Íȵɐ*ǉȾȀ(³ȻǗ%?, @+ÎɯȸɆÉâ?Ņȵɐ*Á)<?8*%?ŅȵɐCȿŲ?ƼɑCŲ"Ĉĥ*ʧÚǾǩ­, ȵɐ*ė%?MZCōť?*+ʋ&$ǅʖȀñÅ@9

8@( y~Lg~

HCƘ?ʔËĈĥ WUSCHEL (WUS)+, ɢˇÎɯȸɆ* organizing center %Ǿǩ, Ņȵɐ*ƜÎãŞCȿŲ?ÁCƘ$ ?(Laux et al., 1996; Mayer et al., 1998)WUS ʧÚǾǩ­%+ưǣyǟǓÞ*ēĐ%8MZōť@, =)˅ǿ )­ȵɐɏ8Íǩ?(Zuo etal., 2002)WUS +ȕʩʇ2 3 Țˏ*MZƫ*ʭ¨ĥǾǩusEH*, 2 "*ƫ%˗

Ǿǩ$ ?(Iwase et al., 2011a)WUS *ƼɑɹƤCʤ7?&%, ɢɥŅȵɐ*ȿŲƼƶ&M
Zōť9Īɏōť*@@*ʋ*ʻʢCǪɹ?&%?%A 

3-4. V bgP_"3&JN.46#u ưǣ*­°>+NCǰ ŉȬ)² =@?(B , ˇȣ)Ħď?ŅȵɐǮ5Í
ȵɐ9*őÎɯ$Ě ?ȵɐC""Ȝ5öB$­°>C$ ?*7, N÷Ĝ*Ÿȋ 4ɫ (@,ǀ ŉȬ+ť>ȡ( ȵɐ÷Ĝ*ŸȋC)Ű!$?*ȵɐěCƶť?\Z, v|\Z, xQa('*ğȯˏ%?, ʛń@=*öť)ʻB?&Ɍ =@?ʰȳʭ¨ĥ*ƼɑƽžĝǶ­%MZōťʏ?&ɲƅĕü@$ ?_nU˚Nicotiana plumbaginifoliat* GLUCURONYL-TRANSFERASE1 (GUT1)*ĝǶ­+ɢɥ*ˇȣ)MZCōť?(Iwai et al., 2002)GUT1 _oQ+ưǣ*xQa*ƶťťÎ*"%? }lN Qch-II )RQʱCʔȘ?ÁCŲ"GUT1 ĝǶ­%+ }lN Qch-II *RQʱw!$ >, ƾȵɐě*{fbQZōť)Ƕłʏ$ ?
 XHjh[h* tumorous shoot development1 (tsd1) & tsd2 ĝǶ­+, ưǣys*
ēĐ%8ȹ£ē˓óɑ(MZCōť?(Frank et al., 2002). TSD1 +ÓRu*ȏȝʧș =
KORRIGAN1 (KOR1) 9 RADIAL SWELLING2 (RSW2)&8ÿ,@$ ?, \Zöť)ʻ
?əȺöđ* endo-1,4-b-D-glucanase CUg$ ?(Nicol et al., 1998; Zuo et al., 2000; Laneet al., 2001; Krupková and Schmülling, 2009)tsd1/kor1/rsw2 ĝǶ­%+, \Zöťǀł)ɫ
B@, Ɩ)xQa*ȸť8ĝã, ȺƦ&$ɢɥ&ƭ*ȸɆã)Ƕłʏ?(Nicol et al.,
1998; His et al., 2001)TSD2 +ÓRu*ɹƤ = QUASIMODO2 (QUA2)9 OVERSENSITIVETO SUGAR1 (OSU1) &CUg$?&Ɍ

ø×%8Ȍ=@$ >, VY­ĸď*~af ZsJ ]=@$?(Mouille et al., 2007; Ralet et al., 2008; Gao et al., 2008)

TSD2/QUA2/OSU1 '*< )ȵɐě*Ǯöť)ʻ$ ? +ƜȌ%?, tsd2/qua2/osu1
ĝǶƫ+xQa*ƶťťÎ*"%?yN Qch 50%8ǔĵ$ ?(Krupková
et al., 2007; Mouille et al., 2007; Ralet et al., 2008)

 tsd1/kor1/rsw2 ĝǶ­*MZōť+, ɢˇÎɯȸɆʻʢĈĥ*ǾǩǶł&WHfMHiś
A. Iwase, M. Ikeuchi & K. Sugimoto-12
BSJ- Revi ew 6: 13 ( 2015)

植物科学最前線Ȩ*¶ʤ)ʏĈ$?

8@((Krupková and Schmülling, 2009)  ²,ƕʠ

SHOOTMERISTEMLESS 9 CLAVATA3 * Ǿ ǩ + ɢ ˇ Î ɯ ȸ Ɇ *  ) Õ ʽ  @ $tsd1/kor1/rsw2 ĝǶ­*MZ%+@=*{Mʭ¨ĥ*Ǿǩɸį@$

6: 14 ( 2015)?,?(Krupková

and Schmülling, 2009)4, tsd1/kor1/rsw2 ĝǶ­%+WHfMHi*śȨƍʤ$ >, W
HfMHiÎɹʰȳ*ʭ¨ĥ%? CYTOKININ OXIDASE1 C tsd1/kor1/rsw2 ĝǶ­%Ǿǩ?&, MZã*ōʎŭÕ@?(Krupková and Schmülling, 2009)@=*ĕü+, ȵɐěťÎ*ǀ öťȸɆ*ȗŇȡ!Îã)Śˉ%>, ÷Ɣ)­ȵɐ*ʧÚ(ĚǃCŭ ?tO)(!$ ?&Cȓ$ ?@=*ȵɐěʻʢĝǶ­%ʏ?ȵɐ*Ěǃ+, ȵɐʺU
|iSX*ƽɤ)<?ʺŸȀ(Ŏˆ 8@( 

3-5. !-% ,*)$?K"3&JN.46#
 DNA *8*9 DNA Ŀ¢pZf_oQʎãħȀ¸˒Cñ?&, DNA ʯÐ*ĝãC©B), Ɣ)£Cʐ $ʭ¨ĥǾǩ*ğƷŞǮ5@?œƣ* DNA ʯÐCʲɷʭ¨ħ)İ$, *< (Õŕ)Ĕ# ʭ¨ħCKrYJkdGQZ&

KrYJkdGbQ(ĝãCʏÕŕĈĥ+, DNA *~aã9pZf*¸˒Cʠ$Q{a*ǥţCĝã, ʔËĈĥ* DNA 1*Ÿɺ*ňö ('Cĝã$ʭ¨ĥǾǩCÕŕ?Kr
YJkdGbQÕŕĈĥ)<?Ġɶƹ(Q{aǥţ*ĝã+ȵɐ*Îã9ɕÎãCUf
?řȀ(ŏÛC$ ?&Ɍ =@$ ?(Gaspar-Maia et al., 2011; Grafi et al., 2011)3
ˏ%+, ǾǮʦĀ*ǌ4!ȵɐ+ʠłQ{aCʸǥţ)$ , Îã&Ç)ǅʖȀĨĪʭ¨ĥǾǩusEH)(!$ *)İ, ğɑŞCŲ"< (ȵɐ+Q{aCʹ ǥţ), `Hh|bQ(ʭ¨ĥǾǩĝã)İ?ǘ¾C$ ? (Gaspar-Maia etal., 2011)
ưǣ%8÷Ʒ*Õŕ? )" $+4ÒǠ&( &Ağ ," *ȵɐħȀ(ȏȝ =, ưǣ*Q{aǥţ8ȵɐ*Îãǥţ)©!$ĝã$ ?&ĕü
@$ ?(Zhao et al., 2001; Verdeil et al., 2007)
Polycomb Repressive Complex1 (PRC1) & PRC2 +ʤãȀ)·Ħ@_oQʎɲö­%>,pZf*ãħȀ¸˒)ʻ$ ?áǣ%+ PRC2 +pZf H3 * 27 ǵȄ*YCf~aã(H3K27me3)?, *
pZf{Q+ B:?ʸQ{aǥţC">, ʭ¨ĥ*ǾǩCŭ ?Ɗ, PRC˜+pZf
 H2A * 119 ǵȄ)?YC l  q O a  ã  ? (H2AK119ub), *pZf{Q8ʛ½)?ʭ¨ĥǾǩ)ŭÕȀ)ÁXIYInK%ǶũȀ(ĆĩōťC?ĝǶ­=Ñ7$

PRC ɵ"=@< ), PRC +Ʒ(ȵɐ*ǾǮʦĀCȿŲ?Á
C?(Ringrose and Paro, 2004)
A. Iwase, M. Ikeuchi & K. Sugimoto-13
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植物科学最前線

ưǣ%+, PRC Îã$ ?Ćĩ%ɏǾǮ&ÎɯȸɆ*uR }Cŭ $ ?, &6: 15 ( 2015)&ğ*²)<!$ȓ@$ ?XHjh[h%+ PRC Cƶť?_oQʎ*ğʲ
ɲ$Ug@$?, @=*ʲĝǶ­, ², PRC2 * CURLY LEAF (CLF) &SWINGER (SWN)*ʲĝǶ­ clf swn 9, VERNALIZATION2 (VRN2)& EMBRYONIC FLOWER2(EMF2)*ʲĝǶ­ emf2 vrn2 +, Ǿɠő48($MZCǮ?(Chanvivattana et al., 2004;
Schubert et al., 2005)  ÷ Ʒ * M  Z ō ť  ¡ * PRC2 * ƶ ť ɴ ȳ *  "FERTILIZATION-INDEPENDENT ENDOSPERM (FIE)*ĝǶ­%8ĕü@$ ?(Bouyer et al.,2011)ưǣ%* PRC1 *Ħď+ʷ ʺȌ=@$ ( !, ăˏ%* RING finger _oQ
ʎ*yR%? At-BMI1A & At-BMI1B ʛń)(!$÷Ī@$ ?(Sanchez-Pulido et al.,2008)PRC2 *ĝǶ­&÷Ʒ), At-bmi1a-1 bmi1b *ʲĝǶ­+Ǿɠőƌ Ǆʿ%MZCōť$4 (Bratzel et al., 2010)
 @=* PRC2 9 PRC1 ĝǶ­*ōʎ+ɏǾǮʻʢĈĥ%? LEC1, LEC2, AGL15, BBM *ǶũȀ(ʧÚǾǩ9 WUS 9 WUSHEL RELATED HOMEOBOX5 (WOX5)('*ŅȵɐʻʢĈĥ*ǶũȀ(Ǿǩ)<!$ŋʏ@$ ?(Bratzel et al., 2010; Bouyer et al., 2011)×ʜ< ), @=*ʭ¨ĥ*3&D'+ʧÚǾǩ%MZǮ?=), @=*ʭ¨ĥ*3&D')+
H3K27me3 9 H2AK119ub *pZf{QÅ!$ ?&ȓ@$ ?"4>@=*ʭ¨ĥ PRC1 9 PRC2 *ȅŸȀ(_Tbf)(!$ >, Ǿǩŭ =@?&%MZãŭ =@$ ?&CŌȓĄ$ ?(Bratzel et al., 2010; Bouyer et al., 2011; Yang et al., 2013)
 PICKLE (PKL) _oQʎ+ Chromodomain-Helicase-DNA binding3 (CHD3)Ru)Îˏ@?Q{aeRsEQ_%>, ʧÚ(ȵɐÎɯCŭ ?*)řȀ(ŏÛCŰ!
$ ?< %?pkl ĝǶ­8Ǿɠő)MZCǮ?(Ogas et al., 1997, 1999)CHD3/
CHD4 Q Z*Q{aeRĈĥ+, áǣ%+pZf*ɕF\aãʰȳ&$Ƽɑ?(Hollender and Liu, 2008)MZʂĳ*Fb\Hȱ%, ĞĈŞ*WHfMHi)İ?
ZzZ!$ ?ĝǶ­&$ cytokinin-hyper-sensitive2 è˃@$ ?, *êĈʭ¨ĥ+ pkl ĝǶ*ÓF%?&Ɨʛ*ȏȝ%Ə= )@$ ?(Furuta et al., 2011)p
ZfɕF\aãʰȳ*ʼĬØ%?fUZ_a A CʳǮƫ)ÌǪ?&WHfMHiśȨ*ƍʤÊǩ@?*%, PKL +pZf*ɕF\aã)Á&ȓĄ@?(Furuta etal., 2011)=) PKL +, H3K27me3 *¸˒)8ʻB!$ ?< %?@+ pkl ĝǶ­%+ LEC1 & LEC2  H3K27me3 {Q*w!$ ?& =Š@$ ?ȺƦ&
$ pkl ĝǶ­%+ LEC1 9 LEC2 Ǿǩ*ŭÕɹʾʏ>MZʂĳ@?(Zhang et al., 2008,2012)
Q{aÕŕĈĥȅŸʔËĈĥ)°ǰQ{aǥţCĝã?&%, ʔËĈĥ*_Tbfʭ¨ĥ*ǾǩCÕŕ?&²ʛńĕü@$?PRC1 *ƶťɴȳ%?

At-BMI1 _  o Q , B3 g ~ H  ʔ Ë Ĉ ĥ %  ? VP1/ABI3-LIKE1 (VAL1; HIGH-LEVELEXPRRESSION OF SUGAR-INDUCIBLE GENE2 [HSI2] &  $ 8 Ȍ = @ $?)&Ⱥö,H2AK119ub C$ LEC1 & LEC2 *ǾǩCŭ $ ?(Yang et al., 2013)4, VAL1/HSI2 *yR%? VAL2/HSI2-LIKE1 (HSL1)+, HISTONE DEACETYLASE19 (HDA19)&Ⱥö, F\aã@$ ?pZf H3(H3Ac)& H4(H4Ac)CɕF\aã?&)<!$ LEC1 &LEC2 *ǾǩCŭ $ ?(Zhou et al., 2013)@=*ĕü¤×) VAL1/ HSI2 & VAL2/HSL1 +,A. Iwase, M. Ikeuchi & K. Sugimoto-14
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植物科学最前線

6: 16 ( 2015)ƼɑʲɲʔËŭÕĈĥ&$, ɠǮ Ɣ)ɏǾǮʻʢĈĥ*ǾǩCŭ ?&%Ʃ˓Ȇ1*
ʔŻC8=$ ?&ĕü@$ (Tsukagoshi et al., 2007)4 hsi2 hsl1 ʲĝǶ­%+ɠǮ ő)MZǮ? (Tsukagoshi et al., 2007)@=CɁö$Ɍ ?&, H2AK119ub 9
H3/H4Ac *ɕF\aã)<!$8Ǿɠő*ȸɆ%*MZãŭÕ@$ ?&Ƞ ?

3-6. :?KUT
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6. I\QY
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